323:
633:. Once the seed is fertilized, the micropyle is sealed by a plug-shaped extension of the nucellus. Unlike living gymnosperms, the tip of the nucellus lacks a pollen chamber (receptacle for stored pollen). The integument is dense and thick, with many layers of differentiated cells. This contrasts with the thin, biseriate (two cell-layer) integument of gnetophytes. Bennettitaleans also lack another gnetophyte-like trait: a sheath of fused bracteoles enveloping the seed. Most integument cells are not unusual in size or shape. However, near the micropyle the innermost layer of integument cells become radially-oriented and elongated, partially closing in on the micropyle. The nucellus and integument are unfused above the
1312:
517:
118:
139:
1278:
1350:
1333:
1260:
336:
1240:
166:
501:
1295:
806:, dubbed the "BEG group". However, this proposal has been contested by other authors, who contend that these similarities are only superficial and do not indicate a close relationship. A 2017 phylogeny based on molecular signatures of fossilised cuticles found that Bennettitales were more closely related to the
798:. Molecular evidence has consistently contradicted the Anthophyte hypothesis, finding that Angiosperms are the sister group to all living gymnosperms, including Gnetales. Some authors have suggested due to similarities between their seed coats, Bennettitales form a clade with the gymnosperm orders of
573:
and fertile sporophylls with ovules at their tips. The presence of ovules at the tips of sporophylls, rather than the tips of stems, is a major difference between the cones of bennettitaleans and gnetophytes. As the cone is fertilized and matures, the microsporophylls wither away and the ovules
298:(cone-bearing seed plants), the relationships of bennettitaleans to other seed plants is debated. Their general resemblance to cycads is contradicted by numerous more subtle features of their reproductive systems and leaf structure. Some authors have linked bennettitaleans to
723:. Buckland provided a description of the family and two species, but failed to give a description of the genus, which has led to Buckland's description of the family being considered invalid by modern taxonomic standards. In publications in 1870, Scottish botanist
692:, providing evidence that this family acted as pollinators for the group. The interseminal scales of Bennettitales ovulate cones may have become fleshy at maturity, which could have potentially made then attractive to wild animals that served as seed dispersers.
640:
Cycadeoidaceans have been suggested to have been self-pollinating, with their stems and cones buried underground, although it has alternatively been proposed that they were pollinated by beetles. The flower-like williamsoniacean male reproductive structure
2123:
Taylor, David
Winship; Li, Hongqi; Dahl, Jeremy; Fago, Fred J.; Zinniker, David; Moldowan, J. Michael (March 2006). "Biogeochemical evidence for the presence of the angiosperm molecular fossil oleanane in Paleozoic and Mesozoic non-angiospermous fossils".
491:
stomata of cycads and conifers. In haplocheilic stomata, the ring of subsidiary cells are not derived from the same original structures as the guard cells. This fundamental difference is the main way to differentiate bennettitalean and cycad foliage.
607:), on the other hand, feature an exposed crown of tapering microsporophylls with adaxial rows of synangia. The microsporophylls may host a single linear row of paired synangia, or instead synangia arranged in a pinnate (feather-shaped) pattern.
284:-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely
923:, and are primarily known from the Cretaceous. Bennettitales were widespread and abundant during the Jurassic and Early Cretaceous, however Bennettitales severely declined during the Late Cretaceous, coincident with the rise of
739:
and the Isle of
Portland. Caruthers was the first to recognise that Bennettitales had distinct differences from cycads, and established the tribes "Williamsonieae" and "Bennettiteae", with the latter being named after the genus
1214:. Most paleobotanists prefer the two families as used here, though some authors, such as Anderson & Anderson (2007), classify the order via a larger number of families. Anderson & Anderson also classified the orders
687:
which had a long, narrow central receptacle which was likely otherwise inaccessible. Early
Cretaceous bennettitalean pollen has been found directly associated with a proboscis bearing fly belonging to the extinct family
470:
The foliage of bennettitaleans resembles that of cycads to such an extent that the foliage of the two groups cannot be reliably distinguished based on gross morphology alone. However, fossil foliage which preserves the
569:, where the pollen and ovules are hosted on the same (bisexual or hermaphrodite) cone. Cavities filled with curved synangia-bearing microsporophylls are encased by thin radiating structures, including thick, infertile
511:
Labels: CFR Centrifugal ray; CPR Centripetal ray; MR Median ridge; FS Fibrous strand; PS Pollen sac position (in this case, pollen sac attachment); CC Central cup; RBA Resin (resinous) body or attractant; ST
1518:"Williamsonia carolinensis sp. nov. and Associated Eoginkgoites Foliage from the Upper Triassic Pekin Formation, North Carolina: Implications for Early Evolution in the Williamsoniaceae (Bennettitales)"
1829:
2554:
2681:
1651:
Osborn, J.M.; Taylor, M.L. (2010). "Pollen and coprolite structure in
Cycadeoidea (Bennettitales): implications for understanding pollination and mating systems in Mesozoic cycadeoids".
2265:"Evolution and Phylogeny of Gnetophytes: Evidence from the Anatomically Preserved Seed Cone Protoephedrites eamesii gen. et sp. nov. and the Seeds of Several Bennettitalean Species"
384:
structures. Williamsoniaceae either had bisporangiate or monosporangiate cones, and distinctly slender and branching woody trunks. The
Williamsoniaceae grew as woody shrubs with a
2408:"Late Palaeozoic Foliage from China Displays Affinities to Cycadales Rather than to Bennettitales Necessitating a Re-Evaluation of the Palaeozoic Pterophyllum Species"
2357:
Wu, Qiong; Ramezani, Jahandar; Zhang, Hua; Wang, Jun; Zeng, Fangui; Zhang, Yichun; Liu, Feng; Chen, Jun; Cai, Yaofeng; Hou, Zhangshuai; Liu, Chao (2021-02-05).
835:
570:
2208:
Friis, Else Marie; Crane, Peter R.; Pedersen, Kaj
Raunsgaard; Bengtson, Stefan; Donoghue, Philip C. J.; Grimm, Guido W.; Stampanoni, Marco (November 2007).
1267:
596:) are similar to mature bisporangiate cones, with interseminal scales and ovule-tipped sporophylls enclosed by bracts. Pollen-producing (male) cones (
2161:"Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms"
1171:
900:) from the Late Triassic of Gondwana appears to be closely related to Bennettitales, but differs from it in some aspects of its reproductive organs.
682:
582:
1259:
1068:
1060:
1016:
546:
surrounded by simple, helically-arranged fertile and infertile structures. Tissue at the base of the cone forms layers of scale-like or petal-like
480:
2678:
2753:
1215:
1139:
1008:
992:
912:
566:
1386:
Peñalver, Enrique; Arillo, Antonio; Pérez-de la Fuente, Ricardo; Riccio, Mark L.; Delclòs, Xavier; Barrón, Eduardo; Grimaldi, David A. (2015).
1311:
1302:
1092:
1052:
818:
488:
443:
322:
1187:
1000:
976:
968:
927:, being mostly extinct by the end of the period, with the final known remains from the Northern Hemisphere being found in the polar latitude
562:. This contrasts with cycads, all of which lack discrete synangia and bear pollen on the abaxial (outer) surface of their microsporophylls.
1099:
1076:
896:
550:
to protect the radiating inner structures. Some authors refer to bennettitalean cones as "flowers", though they are not equivalent to true
907:, and williamsoniaceans became globally distributed by the end of the period. The oldest bennettitalean reproductive structures are small
984:
2359:"High-precision U-Pb age constraints on the Permian floral turnovers, paleoclimate change, and tectonics of the North China block"
1239:
919:. While Williamsoniaceae had a global distribution, Cycadeoidaceae appear to have been primarily confined to the western parts of
407:
In general, bennettitalean leaves are attached to the stem with a helical (corkscrew) arrangement. Some leaves (most species of
399:(not containing all descendants of a common ancestor) assemblage of all Bennettitales that do not belong to the Cycadeoidaceae.
1479:"Anatomically Preserved Early Cretaceous Bennettitalean Leaves: Nilssoniopteris corrugata n. sp. from Vancouver Island, Canada"
2498:"The Ladinian Flora (Middle Triassic) of the Dolomites: palaeoenvironmental reconstructions and palaeoclimatic considerations"
1841:
1744:
Khramov, Alexander V.; Lukashevich, Elena D. (July 2019). "A Jurassic dipteran pollinator with an extremely long proboscis".
1660:
1562:
Blomenkemper, Patrick; Bäumer, Robert; Backer, Malte; Abu Hamad, Abdalla; Wang, Jun; Kerp, Hans; Bomfleur, Benjamin (2021).
1564:"Bennettitalean Leaves From the Permian of Equatorial Pangea—The Early Radiation of an Iconic Mesozoic Gymnosperm Group"
1349:
2827:
868:. True Permian records of benettitalean leaves are rare; outside of the Shihhotse Formation they are only found in the
884:. This formation is notable for the early occurrence of other Mesozoic-style flora, including the earliest records of
2812:
1440:"Divaricate growth habit in Williamsoniaceae (Bennettitales): unravelling the ecology of a key Mesozoic plant group"
585:, with separate pollen and ovule-producing (unisexual) cones on the same plant. The ovule-producing (female) cones (
451:) are incompletely pinnate (sawtooth-shaped) and transitional between these two end members. One unusual leaf form,
2447:
Blomenkemper, Patrick; Kerp, Hans; Abu Hamad, Abdalla; DiMichele, William A.; Bomfleur, Benjamin (December 2018).
680:
produced by bennettitalean reproductive structures, such as the bisexual williamsoniacean reproductive structure
2832:
2310:
Vajda, Vivi; Pucetaite, Milda; McLoughlin, Stephen; Engdahl, Anders; Heimdal, Jimmy; Uvdal, Per (August 2017).
1928:
1897:
862:
do not have conclusive bennettitalean affinities or have been reinterpreted as cycad foliage in the form genus
2137:
2627:
1612:"Is the anthophyte hypothesis alive and well? New evidence from the reproductive structures of Bennettitales"
724:
558:(pollen sacs). The synangia lie on the adaxial (inner) edge of pollen-bearing leaf-like structures known as
1277:
2555:"Ptilophyllum muelleri (Ettingsh.) comb. nov. from the Oligocene of Australia: last of the Bennettitales?"
2497:
1332:
165:
2594:"How similar are the venation and cuticular characters of Glossopteris, Sagenopteris and Anthrophyopsis?"
728:
629:. A long, narrow micropyle extending out of the seed is superficially similar to the condition in living
487:
develop from the same mother cells as the subsidiary cells which surround them. This contrasts with the
2312:"Molecular signatures of fossil leaves provide unexpected new evidence for extinct plant relationships"
637:(base of the seed), unlike cycads or gnetophytes, where the layers are fused for much of their height.
2654:"Heyday of the gymnosperms: systematics and biodiversity of the Late Triassic Molteno fructifications"
2822:
1787:
Peñalver E, Arillo A, Pérez-de la Fuente R, Riccio ML, Delclòs X, Barrón E, Grimaldi DA (July 2015).
932:
617:), with a central embryo surrounded by three layers: the thin megagametophyte, the slightly thicker
2817:
516:
413:, etc.) are narrow, solitary blades with a smooth-edged ("entire") margin. Most leaf morphotypes (
2744:
2690:
2593:
2523:
310:. Molecular data contradicts this, with gnetophytes found to be much more genetically similar to
2807:
2784:
2706:
2653:
2779:
2771:
2210:"Phase-contrast X-ray microtomography links Cretaceous seeds with Gnetales and Bennettitales"
1678:"Diverse sexual strategies in fossil gymnosperms: pollination in the Bennettitales revisited"
285:
117:
2758:
2628:"Brief history of the gymnosperms: classification, biodiversity, phytogeography and ecology"
1517:
1294:
2695:
2460:
2370:
2221:
1753:
1708:
1575:
1163:
1123:
731:
described the first known reproductive organs of the
Bennettitales from Jurassic strata of
648:
587:
522:
415:
328:
148:
8:
751:
543:
381:
130:
2464:
2374:
2225:
1757:
1712:
1579:
2574:
2429:
2388:
2339:
2292:
2264:
2245:
2185:
2160:
2141:
2105:
2093:
2065:
1991:
1944:
1913:
1769:
1726:
1537:
1498:
1459:
1417:
928:
877:
626:
530:
Like other gymnosperms, bennettitalean reproductive inflorescences come in the form of
160:
138:
2766:
2478:
2392:
2331:
2284:
2237:
2190:
2097:
1995:
1837:
1810:
1773:
1656:
1633:
1502:
1478:
1409:
885:
812:
441:(feather-shaped), with many small leaf segments attached to a central shaft. Others (
2578:
2433:
2311:
2296:
2145:
2109:
1730:
1611:
1541:
1463:
1421:
2609:
2605:
2566:
2535:
2468:
2419:
2378:
2343:
2323:
2276:
2249:
2229:
2209:
2180:
2172:
2133:
2089:
2060:
2016:
1983:
1940:
1909:
1875:
1800:
1761:
1716:
1623:
1583:
1529:
1490:
1451:
1399:
1046:
803:
774:
arose from
Bennettitales, as suggested by the wood-like structures and rudimentary
710:
706:
625:. The upper tip of the seed is tapered and opens through a thin and often extended
578:
357:
306:(a rare and unusual group of modern gymnosperms), forming a broader group known as
239:
126:
2539:
1677:
1653:
Plants in deep
Mesozoic time: morphological innovations, phylogeny, and ecosystems
2685:
1198:
1107:
924:
904:
771:
666:
559:
409:
361:
299:
257:
32:
1968:
705:
The
Cycadeoideaceae (originally "Cycadeoideae") were named by English geologist
335:
2035:
Syllabus der Vorlesungen ĂĽber specielle und medicinisch-pharmaceutische Botanik
1024:
353:
234:
190:
153:
2524:"The Late Cretaceous Pterophyllum (Bennettitales) in the North-East of Russia"
2327:
1987:
1805:
1788:
1721:
1696:
1588:
1563:
1494:
1455:
1404:
1387:
2801:
2729:
2288:
2159:
Ran, Jin-Hua; Shen, Ting-Ting; Wang, Ming-Ming; Wang, Xiao-Quan (June 2018).
2010:
1929:"XVII. Contributions towards the History of Zamia Gigas, Lindl. & Hutt.*"
1765:
936:
873:
787:
736:
689:
657:(male and female reproductive structures being present on the same plant) or
472:
261:
68:
2473:
2448:
2020:
1439:
2482:
2335:
2241:
2194:
2176:
2101:
2080:
Crane, P.R. (September 1985). "Phylogenetic relationships in seed plants".
1828:
Friis, Else Marie; Pedersen, Kaj Raunsgaard; Crane, Peter R., eds. (2011),
1814:
1637:
1413:
1357:
1222:
1155:
1131:
1084:
953:
864:
858:), around 281 million years ago. Supposed Carboniferous-Permian records of
778:. Based on morphological data, however, Bennettitales were classified as a
755:
453:
438:
421:
377:
340:
152:
from the Early Cretaceous of India, which may represent an early member of
2553:
McLoughlin, Stephen; Carpenter, Raymond J.; Pott, Christian (April 2011).
2424:
2407:
1864:"The nomenclature of Cycadeoidea (fossil Spermatophyta: Cycadeoideopsida)"
2738:
1969:"The diversity of Australian Mesozoic bennettitopsid reproductive organs"
1340:
1323:
1037:
1030:
742:
719:
672:
Several groups of Jurassic and Early Cretaceous insects possessed a long
43:
2496:
Kustatscher, Evelyn; van Konijnenburg-van Cittert, Johanna H.A. (2005).
2233:
1628:
2495:
1863:
1284:
1147:
1115:
890:
855:
767:
643:
630:
622:
598:
551:
506:
484:
433:
385:
373:
307:
303:
292:
269:
203:
88:
53:
1786:
1385:
2383:
2358:
1898:"XVIII. On Fossil Cycadean Stems from the Secondary Rocks of Britain"
1318:
944:
916:
869:
847:
779:
732:
673:
658:
614:
531:
500:
396:
369:
106:
93:
2700:
2592:
Xu, Yuanyuan; Wang, Yongdong; McLoughlin, Stephen (September 2023).
2048:
1879:
1476:
2723:
2570:
2280:
1533:
1250:
948:
920:
799:
795:
618:
555:
273:
83:
78:
63:
58:
48:
2446:
1967:
McLoughlin, Stephen; Pott, Christian; Sobbe, Ian H. (2018-03-01).
758:
in 1892, who recognised the group as separate from the Cycadales.
2449:"A hidden cradle of plant evolution in Permian tropical lowlands"
1561:
1179:
851:
654:
634:
458:
427:
390:
311:
265:
98:
73:
28:
958:
but no cuticle was preserved, making the referral inconclusive.
272:. Bennettitales were amongst the most common seed plants of the
2626:
Anderson, John M.; Anderson, Heidi M.; Cleal, Chris J. (2007).
2522:
Gnilovskaya, Anastasia A.; Golovneva, Lina B. (February 2018).
2309:
1789:"Long-Proboscid Flies as Pollinators of Cretaceous Gymnosperms"
1610:
Rothwell, Gar W.; Crepet, William L.; Stockey, Ruth A. (2009).
1388:"Long-Proboscid Flies as Pollinators of Cretaceous Gymnosperms"
881:
839:
786:
a study in 2006 suggested that Bennettitales, Angiosperms, and
783:
775:
677:
535:
463:
2406:
Pott, Christian; McLoughlin, Stephen; Lindström, Anna (2009).
830:
The oldest confirmed fossils of bennettitaleans are leaves of
661:(where each plant has only one gender of reproductive organ).
268:
period and became extinct in most areas toward the end of the
1655:. Bloomington, IN: Indiana University Press. pp. 34–49.
843:
791:
770:
hypothesis erected by Arber and Parking in 1907 posited that
714:
547:
539:
476:
365:
281:
277:
177:
1197:
Bennettitales is typically considered the sole order in the
709:
in 1828, from fossil trunks found in Jurassic strata on the
1836:, Cambridge: Cambridge University Press, pp. 101–140,
1477:
Ray, M.M.; Rothwell, G.W.; Stockey, R.A. (September 2014).
610:
2207:
2138:
10.1666/0094-8373(2006)32[179:BEFTPO]2.0.CO;2
810:+Cycads clade than conifers, and were closely related to
754:. The order Bennettitales was erected by German botanist
314:. The exact position of Bennettitales remains uncertain.
1650:
943:
A possible late record has been reported from the early
653:, though it is uncertain whether the parent plants were
520:
Cross section of the female williamsoniaceous seed cone
504:
Diagram of male Williamsoniaceae reproductive structure
2552:
2405:
1830:"Angiosperms in context: extant and fossil seed plants"
2625:
1609:
761:
647:
is associated with the female reproductive structure
542:(unfertilized seeds). The cones have a thick central
475:
can be assigned to either group with confidence. The
2521:
1516:
Pott, Christian; Axsmith, Brian J. (February 2015).
669:, with some species possibly pollinated by beetles.
2356:
2012:
A biographical index of British and Irish botanists
1966:
1676:Frame, Dawn; Gottsberger, Gerhard (21 April 2023).
1438:Pott, Christian; McLoughlin, Stephen (2014-06-01).
903:The bennettitalean fossil record reappeared in the
1827:
1743:
395:It has been suggested that Williamsoniaceae are a
2263:Rothwell, Gar W.; Stockey, Ruth A. (March 2013).
894:). The order Fredlindiales (containing the genus
368:-like trunks with bisporangiate (containing both
2799:
2691:Lecture Notes on Mesozoic Mesophytic Communities
2651:
2591:
2122:
2009:Britten, James; Boulger, George Simonds (1893).
1675:
2262:
2158:
1697:"Review of the bennettitalean genus Weltrichia"
2652:Anderson, John M.; Anderson, Heidi M. (2003).
2008:
1437:
2015:. London: West, Newman & co. p. 14.
1933:Transactions of the Linnean Society of London
1902:Transactions of the Linnean Society of London
834:, a species from the upper part of the Upper
750:the name being in honour of British botanist
676:, and it has been suggested that they fed on
352:Bennettitales are divided into two families,
2696:Weblinks to articles about the Bennettitales
554:flowers. Pollen is often enclosed in paired
326:Bennettitales foliage assigned to the genus
2046:
2037:. xxiii + 184 pp. Gebr. Borntraeger, Berlin
1515:
1926:
1895:
746:named by Caruthers in the same publication
137:
116:
2472:
2423:
2382:
2184:
2064:
1976:Palaeobiodiversity and Palaeoenvironments
1804:
1720:
1627:
1587:
1444:Palaeobiodiversity and Palaeoenvironments
1403:
339:Restoration of the willamsoniacean shrub
2053:Botanical Journal of the Linnean Society
515:
499:
334:
321:
2559:International Journal of Plant Sciences
2269:International Journal of Plant Sciences
1522:International Journal of Plant Sciences
825:
700:
461:appearance similar to early species of
2800:
2621:
2619:
1861:
1834:Early Flowers and Angiosperm Evolution
972:(scales or bracts, unplaced in family)
735:and Jurassic-Cretaceous strata of the
2705:
2704:
2598:Review of Palaeobotany and Palynology
2079:
2047:Arber, E.N.; Parkin, J. (July 1907).
1962:
1960:
1958:
1956:
1954:
1891:
1889:
1694:
1605:
1603:
1601:
1599:
1557:
1555:
1553:
1551:
1433:
1431:
1381:
1379:
1377:
1375:
846:. This strata is dated to the early
479:of bennettitaleans are described as
388:branching habit, similar to that of
2616:
1862:Doweld, Alexander B. (2016-05-03).
1012:(ovulate organ, unplaced in family)
911:"flowers" from the Middle Triassic
713:, England, which Buckland gave the
577:Most bennettitaleans in the family
13:
2546:
2440:
2201:
2094:10.1111/j.1096-0031.1985.tb00432.x
2066:10.1111/j.1095-8339.1907.tb01074.x
1951:
1945:10.1111/j.1096-3642.1870.tb00200.x
1914:10.1111/j.1096-3642.1870.tb00201.x
1886:
1780:
1695:Popa, Mihai Emilian (2019-02-27).
988:(pollen organ, unplaced in family)
762:Relationships to other seed plants
495:
483:. This means that the main paired
348:showing divaricate branching habit
14:
2844:
2672:
1596:
1548:
1428:
1372:
1175:(bisexual reproductive structure)
276:, and had morphologies including
2165:Proceedings. Biological Sciences
1896:Carruthers, William (May 1870).
1348:
1331:
1310:
1293:
1276:
1258:
1238:
613:are dicotyledonous (possess two
164:
41:
2645:
2585:
2515:
2489:
2399:
2350:
2303:
2256:
2152:
2116:
2073:
2040:
2027:
2002:
1920:
1855:
1821:
1737:
2610:10.1016/j.revpalbo.2023.104934
2316:Nature Ecology & Evolution
2049:"On the Origin of Angiosperms"
1927:Williamson, W. C. (May 1870).
1688:
1669:
1644:
1509:
1470:
1219:Anderson & Anderson (2003)
317:
1:
2540:10.1016/j.cretres.2017.09.013
2412:Acta Palaeontologica Polonica
1365:
1226:Pilger & Melchior (1954)
961:
364:. Cycadeoidaceae had stout,
16:Extinct order of seed plants
7:
1271:from the Jurassic of Mexico
832:Nilssoniopteris shanxiensis
729:William Crawford Williamson
695:
264:that first appeared in the
125:Restoration of a member of
10:
2849:
1701:Journal of Palaeogeography
1616:American Journal of Botany
1568:Frontiers in Earth Science
1231:
951:, assignable to the genus
727:and English paleobotanist
402:
2828:Permian first appearances
2713:
2328:10.1038/s41559-017-0224-5
1988:10.1007/s12549-017-0286-z
1806:10.1016/j.cub.2015.05.062
1722:10.1186/s42501-019-0023-9
1589:10.3389/feart.2021.652699
1495:10.1017/S002233600005767X
1456:10.1007/s12549-014-0157-9
1405:10.1016/j.cub.2015.05.062
1358:Ptilophyllum grandifolium
947:of eastern Australia and
794:based on the presence of
565:Many bennettitaleans are
380:(cones) serving as their
231:
226:
161:Scientific classification
159:
149:"Williamsonia" sewardiana
145:
136:
124:
115:
23:
2813:Prehistoric plant orders
1766:10.1016/j.gr.2019.02.004
1228:within Bennettitopsida.
935:, Russia, dating to the
39:Kungurian –Maastrichtian
2474:10.1126/science.aau4061
2021:10.5962/bhl.title.56947
1483:Journal of Paleontology
302:(flowering plants) and
2177:10.1098/rspb.2018.1012
574:transform into seeds.
527:
513:
360:, which have distinct
349:
332:
2833:Oligocene extinctions
2780:Paleobiology Database
2425:10.4202/app.2009.0070
2033:Engler, H.G.A. 1892.
665:was likely primarily
621:, and the protective
519:
503:
338:
325:
286:pollinated by insects
826:Evolutionary history
701:History of discovery
457:, even approaches a
146:Life restoration of
2528:Cretaceous Research
2465:2018Sci...362.1414B
2459:(6421): 1414–1416.
2375:2021Geo....49..677W
2234:10.1038/nature06278
2226:2007Natur.450..549F
1758:2019GondR..71..210K
1713:2019JPalg...8...12P
1629:10.3732/ajb.0800209
1580:2021FrEaS...9..162B
1167:(female seed cones)
1151:(male pollen organ)
1072:(male pollen organ)
850:stage of the early
836:Shihhotse Formation
752:John Joseph Bennett
571:interseminal scales
447:, a few species of
291:Although certainly
2684:2015-04-02 at the
2171:(1881): 20181012.
1247:Zamites mariposana
1143:(female seed cone)
1064:(female seed cone)
929:Kakanaut Formation
878:Umm Irna Formation
780:monophyletic group
725:William Carruthers
528:
514:
350:
333:
2795:
2794:
2767:Open Tree of Life
2707:Taxon identifiers
1843:978-0-521-59283-3
1746:Gondwana Research
1662:978-0-253-00199-3
1398:(14): 1917–1923.
1227:
1220:
1213:
1206:
1183:(leaf, in partim)
1004:? (pollen organ?)
996:? (pollen organ?)
980:? (pollen organ?)
886:corystospermalean
788:Gigantopteridales
782:when paired with
247:
246:
222:
110:
2840:
2823:Paleogene plants
2788:
2787:
2775:
2774:
2762:
2761:
2749:
2748:
2747:
2734:
2733:
2732:
2702:
2701:
2666:
2665:
2649:
2643:
2642:
2632:
2623:
2614:
2613:
2589:
2583:
2582:
2550:
2544:
2543:
2519:
2513:
2512:
2502:
2493:
2487:
2486:
2476:
2444:
2438:
2437:
2427:
2403:
2397:
2396:
2386:
2384:10.1130/G48051.1
2354:
2348:
2347:
2322:(8): 1093–1099.
2307:
2301:
2300:
2260:
2254:
2253:
2220:(7169): 549–52.
2205:
2199:
2198:
2188:
2156:
2150:
2149:
2120:
2114:
2113:
2077:
2071:
2070:
2068:
2044:
2038:
2031:
2025:
2024:
2006:
2000:
1999:
1973:
1964:
1949:
1948:
1924:
1918:
1917:
1893:
1884:
1883:
1859:
1853:
1852:
1851:
1850:
1825:
1819:
1818:
1808:
1784:
1778:
1777:
1741:
1735:
1734:
1724:
1692:
1686:
1685:
1673:
1667:
1666:
1648:
1642:
1641:
1631:
1607:
1594:
1593:
1591:
1559:
1546:
1545:
1513:
1507:
1506:
1474:
1468:
1467:
1435:
1426:
1425:
1407:
1383:
1352:
1335:
1314:
1297:
1280:
1268:Weltrichia magna
1262:
1242:
1225:
1218:
1211:
1209:Cycadeoideopsida
1204:
1047:Williamsoniaceae
939:, assignable to
925:flowering plants
804:Erdtmanithecales
711:Isle of Portland
707:William Buckland
615:embryonic leaves
579:Williamsoniaceae
560:microsporophylls
534:, which produce
358:Williamsoniaceae
256:) is an extinct
240:Williamsoniaceae
220:
215:
169:
168:
141:
127:Williamsoniaceae
120:
104:
103:
40:
27:Temporal range:
21:
20:
2848:
2847:
2843:
2842:
2841:
2839:
2838:
2837:
2818:Mesozoic plants
2798:
2797:
2796:
2791:
2783:
2778:
2770:
2765:
2757:
2752:
2743:
2742:
2737:
2728:
2727:
2722:
2709:
2686:Wayback Machine
2675:
2670:
2669:
2650:
2646:
2630:
2624:
2617:
2590:
2586:
2551:
2547:
2520:
2516:
2500:
2494:
2490:
2445:
2441:
2404:
2400:
2355:
2351:
2308:
2304:
2261:
2257:
2206:
2202:
2157:
2153:
2121:
2117:
2078:
2074:
2045:
2041:
2032:
2028:
2007:
2003:
1971:
1965:
1952:
1925:
1921:
1894:
1887:
1880:10.12705/652.16
1860:
1856:
1848:
1846:
1844:
1826:
1822:
1799:(14): 1917–23.
1793:Current Biology
1785:
1781:
1742:
1738:
1693:
1689:
1674:
1670:
1663:
1649:
1645:
1608:
1597:
1560:
1549:
1514:
1510:
1475:
1471:
1436:
1429:
1392:Current Biology
1384:
1373:
1368:
1361:
1353:
1344:
1336:
1327:
1315:
1306:
1298:
1289:
1281:
1272:
1265:Restoration of
1263:
1254:
1245:Fossil leaf of
1243:
1234:
1202:Bennettitopsida
1172:Williamsoniella
1108:Nilssoniopteris
964:
905:Middle Triassic
840:Shanxi Province
828:
764:
703:
698:
683:Williamsoniella
667:wind-pollinated
583:monosporangiate
498:
496:Cones and seeds
449:Nilssoniopteris
410:Nilssoniopteris
405:
320:
252:(also known as
219:
213:
163:
111:
102:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
38:
37:
36:
33:Late Cretaceous
25:
17:
12:
11:
5:
2846:
2836:
2835:
2830:
2825:
2820:
2815:
2810:
2793:
2792:
2790:
2789:
2776:
2763:
2750:
2735:
2719:
2717:
2711:
2710:
2699:
2698:
2693:
2688:
2674:
2673:External links
2671:
2668:
2667:
2644:
2615:
2584:
2571:10.1086/658920
2545:
2514:
2488:
2439:
2418:(1): 157–168.
2398:
2369:(6): 677–681.
2349:
2302:
2281:10.1086/668688
2275:(3): 511–529.
2255:
2200:
2151:
2115:
2088:(4): 329–348.
2072:
2059:(263): 29–80.
2039:
2026:
2001:
1950:
1939:(4): 663–674.
1919:
1908:(4): 675–708.
1885:
1874:(2): 372–379.
1854:
1842:
1820:
1779:
1736:
1687:
1668:
1661:
1643:
1622:(1): 296–322.
1595:
1547:
1534:10.1086/679471
1528:(2): 174–185.
1508:
1489:(5): 1085–93.
1469:
1450:(2): 307–325.
1427:
1370:
1369:
1367:
1364:
1363:
1362:
1354:
1347:
1345:
1337:
1330:
1328:
1316:
1309:
1307:
1299:
1292:
1290:
1282:
1275:
1273:
1264:
1257:
1255:
1244:
1237:
1233:
1230:
1195:
1194:
1193:
1192:
1191:(pollen organ)
1184:
1176:
1168:
1160:
1152:
1144:
1136:
1128:
1120:
1112:
1104:
1096:
1089:
1081:
1073:
1069:Bennettistemon
1065:
1061:Bennetticarpus
1057:
1043:
1042:
1041:
1034:
1025:Cycadeoidaceae
1021:
1017:Anthrophyopsis
1013:
1005:
997:
989:
981:
973:
963:
960:
827:
824:
763:
760:
702:
699:
697:
694:
497:
494:
481:syndetocheilic
404:
401:
354:Cycadeoidaceae
319:
316:
245:
244:
243:
242:
237:
235:Cycadeoidaceae
229:
228:
224:
223:
211:
207:
206:
204:Spermatophytes
201:
194:
193:
188:
181:
180:
175:
171:
170:
157:
156:
154:Cycadeoidaceae
143:
142:
134:
133:
131:Thérèse Ekblom
122:
121:
113:
112:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
26:
15:
9:
6:
4:
3:
2:
2845:
2834:
2831:
2829:
2826:
2824:
2821:
2819:
2816:
2814:
2811:
2809:
2808:Bennettitales
2806:
2805:
2803:
2786:
2781:
2777:
2773:
2768:
2764:
2760:
2755:
2751:
2746:
2745:Bennettitales
2740:
2736:
2731:
2725:
2721:
2720:
2718:
2716:
2715:Bennettitales
2712:
2708:
2703:
2697:
2694:
2692:
2689:
2687:
2683:
2680:
2677:
2676:
2663:
2659:
2655:
2648:
2640:
2636:
2629:
2622:
2620:
2611:
2607:
2603:
2599:
2595:
2588:
2580:
2576:
2572:
2568:
2565:(4): 574–85.
2564:
2560:
2556:
2549:
2541:
2537:
2533:
2529:
2525:
2518:
2510:
2506:
2499:
2492:
2484:
2480:
2475:
2470:
2466:
2462:
2458:
2454:
2450:
2443:
2435:
2431:
2426:
2421:
2417:
2413:
2409:
2402:
2394:
2390:
2385:
2380:
2376:
2372:
2368:
2364:
2360:
2353:
2345:
2341:
2337:
2333:
2329:
2325:
2321:
2317:
2313:
2306:
2298:
2294:
2290:
2286:
2282:
2278:
2274:
2270:
2266:
2259:
2251:
2247:
2243:
2239:
2235:
2231:
2227:
2223:
2219:
2215:
2211:
2204:
2196:
2192:
2187:
2182:
2178:
2174:
2170:
2166:
2162:
2155:
2147:
2143:
2139:
2135:
2132:(2): 179–90.
2131:
2127:
2119:
2111:
2107:
2103:
2099:
2095:
2091:
2087:
2083:
2076:
2067:
2062:
2058:
2054:
2050:
2043:
2036:
2030:
2022:
2018:
2014:
2013:
2005:
1997:
1993:
1989:
1985:
1981:
1977:
1970:
1963:
1961:
1959:
1957:
1955:
1946:
1942:
1938:
1934:
1930:
1923:
1915:
1911:
1907:
1903:
1899:
1892:
1890:
1881:
1877:
1873:
1869:
1865:
1858:
1845:
1839:
1835:
1831:
1824:
1816:
1812:
1807:
1802:
1798:
1794:
1790:
1783:
1775:
1771:
1767:
1763:
1759:
1755:
1751:
1747:
1740:
1732:
1728:
1723:
1718:
1714:
1710:
1706:
1702:
1698:
1691:
1683:
1679:
1672:
1664:
1658:
1654:
1647:
1639:
1635:
1630:
1625:
1621:
1617:
1613:
1606:
1604:
1602:
1600:
1590:
1585:
1581:
1577:
1573:
1569:
1565:
1558:
1556:
1554:
1552:
1543:
1539:
1535:
1531:
1527:
1523:
1519:
1512:
1504:
1500:
1496:
1492:
1488:
1484:
1480:
1473:
1465:
1461:
1457:
1453:
1449:
1445:
1441:
1434:
1432:
1423:
1419:
1415:
1411:
1406:
1401:
1397:
1393:
1389:
1382:
1380:
1378:
1376:
1371:
1360:
1359:
1351:
1346:
1343:
1342:
1334:
1329:
1326:
1325:
1320:
1317:Seed bearing
1313:
1308:
1305:
1304:
1296:
1291:
1287:
1286:
1279:
1274:
1270:
1269:
1261:
1256:
1252:
1248:
1241:
1236:
1235:
1229:
1224:
1217:
1216:Fredlindiales
1210:
1205:Engler (1897)
1203:
1200:
1190:
1189:
1185:
1182:
1181:
1177:
1174:
1173:
1169:
1166:
1165:
1161:
1159:(whole plant)
1158:
1157:
1153:
1150:
1149:
1145:
1142:
1141:
1140:Vardekloeftia
1137:
1134:
1133:
1129:
1126:
1125:
1121:
1118:
1117:
1113:
1110:
1109:
1105:
1103:(whole plant)
1102:
1101:
1097:
1095:
1094:
1090:
1087:
1086:
1082:
1079:
1078:
1074:
1071:
1070:
1066:
1063:
1062:
1058:
1055:
1054:
1050:
1049:
1048:
1044:
1040:
1039:
1035:
1033:
1032:
1028:
1027:
1026:
1022:
1019:
1018:
1014:
1011:
1010:
1009:Westersheimia
1006:
1003:
1002:
998:
995:
994:
993:Leguminanthus
990:
987:
986:
982:
979:
978:
974:
971:
970:
966:
965:
959:
957:
955:
950:
946:
942:
941:Pterophyllum.
938:
937:Maastrichtian
934:
930:
926:
922:
918:
914:
913:Esk Formation
910:
906:
901:
899:
898:
893:
892:
887:
883:
879:
875:
874:Changhsingian
871:
867:
866:
861:
857:
853:
849:
845:
841:
837:
833:
823:
822:
820:
815:
814:
809:
805:
801:
797:
793:
789:
785:
781:
777:
773:
769:
759:
757:
753:
749:
745:
744:
738:
737:Isle of Wight
734:
730:
726:
722:
721:
716:
712:
708:
693:
691:
690:Zhangsolvidae
686:
684:
679:
675:
670:
668:
664:
660:
656:
652:
651:
646:
645:
638:
636:
632:
628:
624:
620:
616:
612:
608:
606:
602:
600:
595:
591:
589:
584:
580:
575:
572:
568:
567:bisporangiate
563:
561:
557:
553:
549:
545:
541:
537:
533:
526:
524:
518:
510:
508:
502:
493:
490:
486:
482:
478:
474:
468:
466:
465:
460:
456:
455:
450:
446:
445:
440:
436:
435:
430:
429:
424:
423:
418:
417:
412:
411:
400:
398:
394:
392:
387:
383:
379:
375:
371:
367:
363:
362:growth habits
359:
355:
347:
346:
343:
337:
331:
330:
324:
315:
313:
309:
305:
301:
297:
294:
289:
287:
283:
279:
275:
271:
267:
263:
259:
255:
251:
250:Bennettitales
241:
238:
236:
233:
232:
230:
225:
218:
217:Bennettitales
212:
209:
208:
205:
202:
199:
196:
195:
192:
191:Tracheophytes
189:
186:
183:
182:
179:
176:
173:
172:
167:
162:
158:
155:
151:
150:
144:
140:
135:
132:
128:
123:
119:
114:
108:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
34:
30:
24:Bennettitales
22:
19:
2714:
2661:
2657:
2647:
2638:
2634:
2601:
2597:
2587:
2562:
2558:
2548:
2531:
2527:
2517:
2508:
2504:
2491:
2456:
2452:
2442:
2415:
2411:
2401:
2366:
2362:
2352:
2319:
2315:
2305:
2272:
2268:
2258:
2217:
2213:
2203:
2168:
2164:
2154:
2129:
2126:Paleobiology
2125:
2118:
2085:
2081:
2075:
2056:
2052:
2042:
2034:
2029:
2011:
2004:
1982:(1): 71–95.
1979:
1975:
1936:
1932:
1922:
1905:
1901:
1871:
1867:
1857:
1847:, retrieved
1833:
1823:
1796:
1792:
1782:
1749:
1745:
1739:
1704:
1700:
1690:
1681:
1671:
1652:
1646:
1619:
1615:
1571:
1567:
1525:
1521:
1511:
1486:
1482:
1472:
1447:
1443:
1395:
1391:
1356:
1339:
1322:
1303:Anomozamites
1301:
1283:
1266:
1246:
1223:Pentoxylales
1212:Scott (1923)
1208:
1201:
1196:
1186:
1178:
1170:
1164:Williamsonia
1162:
1156:Wielandiella
1154:
1146:
1138:
1132:Ptilophyllum
1130:
1124:Pterophyllum
1122:
1114:
1106:
1098:
1093:Ischnophyton
1091:
1085:Eoginkgoites
1083:
1075:
1067:
1059:
1053:Anomozamites
1051:
1036:
1029:
1015:
1007:
999:
991:
983:
975:
967:
954:Ptilophyllum
952:
940:
909:Williamsonia
908:
902:
895:
889:
870:Late Permian
865:Pseudoctenis
863:
860:Pterophyllum
859:
831:
829:
819:Ptilozamites
817:
811:
807:
765:
756:Adolf Engler
747:
741:
718:
704:
681:
671:
662:
650:Williamsonia
649:
642:
639:
609:
604:
597:
593:
588:Williamsonia
586:
581:are instead
576:
564:
529:
523:Williamsonia
521:
505:
489:haplocheilic
469:
462:
454:Eoginkgoites
452:
448:
444:Anomozamites
442:
437:, etc.) are
432:
426:
422:Ptilophyllum
420:
416:Pterophyllum
414:
408:
406:
397:paraphyletic
389:
382:reproductive
351:
345:angustifolia
344:
342:Wielandiella
341:
329:Pterophyllum
327:
295:
290:
253:
249:
248:
221:Engler, 1892
216:
197:
184:
147:
18:
2739:Wikispecies
2679:Gymnosperms
1752:: 210–215.
1341:Cycadeoidea
1324:Bennettites
1188:Haitingeria
1038:Monanthesia
1031:Cycadeoidea
1001:Leuthardtia
977:Haitingeria
969:Cycadolepis
772:angiosperms
743:Bennettites
720:Cycadeoidea
631:gnetophytes
485:guard cells
374:microspores
318:Description
304:gnetophytes
300:angiosperms
293:gymnosperms
262:seed plants
2802:Categories
2658:Strelitzia
2635:Strelitzia
2604:: 104934.
2082:Cladistics
1849:2023-03-20
1366:References
1355:Fossil of
1285:Otozamites
1148:Weltrichia
1116:Otozamites
1100:Kimuriella
1077:Bucklandia
897:Fredlindia
891:Dicroidium
856:Cisuralian
768:Anthophyte
663:Weltrichia
655:monoecious
644:Weltrichia
623:integument
599:Weltrichia
552:angiosperm
544:receptacle
525:harrisiana
507:Weltrichia
434:Otozamites
386:divaricate
370:megaspores
308:Anthophyta
296:sensu lato
270:Cretaceous
254:cycadeoids
2534:: 56–63.
2393:234064909
2289:1058-5893
1996:135237376
1774:134847380
1707:(1): 12.
1503:232349931
1338:Trunk of
1319:strobilus
1249:from the
962:Subgroups
945:Oligocene
917:Australia
888:foliage (
848:Kungurian
813:Nilssonia
784:Gnetales.
733:Yorkshire
674:proboscis
659:dioecious
627:micropyle
227:Families
174:Kingdom:
107:Oligocene
105:Possible
2724:Wikidata
2682:Archived
2664:: 1–308.
2641:: 1–280.
2579:52885618
2511:: 31–51.
2483:30573628
2434:52253114
2336:29046567
2297:84063572
2242:18033296
2195:29925623
2146:83801635
2110:85709030
2102:34965681
1815:26166781
1731:85458053
1684:. 62/63.
1638:21628190
1542:44559347
1464:84440045
1422:13022302
1414:26166781
1300:Leaf of
1251:Jurassic
1020:? (leaf)
949:Tasmania
933:Chukotka
921:Laurasia
872:(likely
800:Gnetales
796:oleanane
696:Taxonomy
619:nucellus
556:synangia
378:strobili
312:conifers
274:Mesozoic
2772:5308459
2730:Q132823
2505:Geo.Alp
2461:Bibcode
2453:Science
2371:Bibcode
2363:Geology
2344:3604369
2250:1198220
2222:Bibcode
2186:6030518
1754:Bibcode
1709:Bibcode
1576:Bibcode
1574:: 162.
1232:Gallery
1180:Zamites
1045:Family
1023:Family
852:Permian
790:form a
776:flowers
635:chalaza
477:stomata
473:cuticle
459:palmate
439:pinnate
428:Zamites
403:Foliage
391:Banksia
266:Permian
210:Order:
178:Plantae
29:Permian
2577:
2481:
2432:
2391:
2342:
2334:
2295:
2287:
2248:
2240:
2214:Nature
2193:
2183:
2144:
2108:
2100:
1994:
1840:
1813:
1772:
1729:
1682:Phyton
1659:
1636:
1540:
1501:
1462:
1420:
1412:
1135:(leaf)
1127:(leaf)
1119:(leaf)
1111:(leaf)
1088:(leaf)
1080:(axes)
1056:(leaf)
985:Lunzia
882:Jordan
876:)-age
808:Ginkgo
678:nectar
548:bracts
540:ovules
536:pollen
464:Ginkgo
109:record
2785:55214
2759:11050
2754:IRMNG
2631:(PDF)
2575:S2CID
2501:(PDF)
2430:S2CID
2389:S2CID
2340:S2CID
2293:S2CID
2246:S2CID
2142:S2CID
2106:S2CID
1992:S2CID
1972:(PDF)
1868:Taxon
1770:S2CID
1727:S2CID
1538:S2CID
1499:S2CID
1460:S2CID
1418:S2CID
1199:class
844:China
792:clade
717:name
715:genus
611:Seeds
532:cones
512:Stalk
366:cycad
282:cycad
278:shrub
258:order
198:Clade
185:Clade
2479:PMID
2332:PMID
2285:ISSN
2238:PMID
2191:PMID
2098:PMID
1838:ISBN
1811:PMID
1657:ISBN
1634:PMID
1410:PMID
1288:leaf
1221:and
816:and
802:and
766:The
538:and
372:and
356:and
280:and
44:Preęž’
2606:doi
2602:316
2567:doi
2563:172
2536:doi
2469:doi
2457:362
2420:doi
2379:doi
2324:doi
2277:doi
2273:174
2230:doi
2218:450
2181:PMC
2173:doi
2169:285
2134:doi
2090:doi
2061:doi
2017:doi
1984:doi
1941:doi
1910:doi
1876:doi
1801:doi
1762:doi
1717:doi
1624:doi
1584:doi
1530:doi
1526:176
1491:doi
1452:doi
1400:doi
1321:of
1207:or
931:in
915:of
880:in
838:in
603:etc
592:etc
260:of
129:by
2804::
2782::
2769::
2756::
2741::
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2662:15
2660:.
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2637:.
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